HANDBOOK of FISH BIOLOGY and FISHERIES Volume 1 Also Available from Blackwell Publishing: Handbook of Fish Biology and Fisheries Edited by Paul J.B
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HANDBOOK OF FISH BIOLOGY AND FISHERIES Volume 1 Also available from Blackwell Publishing: Handbook of Fish Biology and Fisheries Edited by Paul J.B. Hart and John D. Reynolds Volume 2 Fisheries Handbook of Fish Biology and Fisheries VOLUME 1 FISH BIOLOGY EDITED BY Paul J.B. Hart Department of Biology University of Leicester AND John D. Reynolds School of Biological Sciences University of East Anglia © 2002 by Blackwell Science Ltd a Blackwell Publishing company Chapter 8 © British Crown copyright, 1999 BLACKWELL PUBLISHING 350 Main Street, Malden, MA 02148‐5020, USA 108 Cowley Road, Oxford OX4 1JF, UK 550 Swanston Street, Carlton, Victoria 3053, Australia The right of Paul J.B. Hart and John D. Reynolds to be identified as the Authors of the Editorial Material in this Work has been asserted in accordance with the UK Copyright, Designs, and Patents Act 1988. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs, and Patents Act 1988, without the prior permission of the publisher. First published 2002 Reprinted 2004 Library of Congress Cataloging‐in‐Publication Data has been applied for. Volume 1 ISBN 0‐632‐05412‐3 (hbk) Volume 2 ISBN 0‐632‐06482‐X (hbk) 2‐volume set ISBN 0‐632‐06483‐8 A catalogue record for this title is available from the British Library. Set in 9/11.5 pt Trump Mediaeval by SNP Best‐set Typesetter Ltd, Hong Kong Printed and bound in the United Kingdom by TJ International Ltd, Padstow, Cornwall. The publisher’s policy is to use permanent paper from mills that operate a sustainable forestry policy, and which has been manufactured from pulp processed using acid‐free and elementary chlorine‐free practices. Furthermore, the publisher ensures that the text paper and cover board used have met acceptable environmental accreditation standards. For further information on Blackwell Publishing, visit our website: http://www.blackwellpublishing.com Contents List of Contributors, x Preface, xii List of Abbreviations, xiv 1 BANISHING IGNORANCE: UNDERPINNING FISHERIES WITH BASIC BIOLOGY, 1 Paul J.B. Hart and John D. Reynolds 1.1 Introduction, 1 1.2 Global fisheries, 1 1.3 The quest for knowledge, 3 1.4 Part 1: Biodiversity, 4 1.5 Part 2: Production and population structure, 5 1.6 Part 3: Fish as predators and prey, 7 1.7 Part 4: Fish in ecosystems, 8 1.8 Ignorance banished?, 9 1.9 Conclusions, 10 Part 1: Biodiversity, 13 2 PHYLOGENY AND SYSTEMATICS OF FISHES, 15 A.C. Gill and R.D. Mooi 2.1 Introduction, 15 2.2 Phylogenetic methods and classification, 15 2.3 Fish diversity and phylogeny, 20 2.4 Conclusions, 36 3 HISTORICAL BIOGEOGRAPHY OF FISHES, 43 R.D. Mooi and A.C. Gill 3.1 Introduction, 43 3.2 Concepts and methods, 44 3.3 Distribution, faunal composition and historical biogeography by region, 47 3.4 Conclusions, 62 vi Contents Part 2: Production and Population Structure, 69 4 THE PHYSIOLOGY OF LIVING IN WATER, 71 Ole Brix 4.1 Introduction, 71 4.2 Buoyancy, or coping with pressure, 72 4.3 Swimming, 75 4.4 Osmoregulatory problems in fresh and salt water, 78 4.5 Respiration and special adaptations for living in low oxygen, 82 4.6 Digestion and absorption, 90 4.7 Bioluminescence, 91 4.8 Conclusions, 92 5 ENVIRONMENTAL FACTORS AND RATES OF DEVELOPMENT AND GROWTH, 97 Malcolm Jobling 5.1 Introduction, 97 5.2 Terminology of life-history stages, 97 5.3 Development and growth during early life history, 99 5.4 Growth models and equations, 102 5.5 Age determination, back-calculation and validation techniques, 104 5.6 Length–weight relationships and indices of condition and growth, 107 5.7 Energy budget and bioenergetics: energy partitioning and storage, 109 5.8 Growth at different latitudes: models of growth compensation, 113 5.9 Estimating food consumption, 115 5.10 Conclusions, 117 6 RECRUITMENT: UNDERSTANDING DENSITY-DEPENDENCE IN FISH POPULATIONS, 123 Ransom A. Myers 6.1 Introduction, 123 6.2 The link between spawner abundance and subsequent recruitment, 124 6.3 Generalities through meta-analysis, 129 6.4 Carrying capacity, 130 6.5 Variability in recruitment, 131 6.6 At what life-history stage does density-dependent mortality occur?, 131 6.7 Estimating density-dependent mortality from long-term surveys, 133 6.8 Pelagic egg, larval and juvenile stages, 136 6.9 Future research, 141 6.10 Conclusions, 144 Contents vii 7 LIFE HISTORIES OF FISH, 149 J.A. Hutchings 7.1 Introduction, 149 7.2 Influence of survival and growth rate on age, size and reproductive effort at maturity, 152 7.3 Offspring size and number strategies ,158 7.4 Alternative life-history strategies, 162 7.5 Effects of fishing on life history, 165 7.6 Conclusions, 167 8 MIGRATION, 175 Julian Metcalfe, Geoff Arnold and Robert McDowall 8.1 Introduction, 175 8.2 Exploitation and ecology, 178 8.3 Fish migrations, 179 8.4 Migratory mechanisms, 189 8.5 Techniques, 191 8.6 Distribution and genetics, 192 8.7 Fishery applications, 194 8.8 Conclusions, 194 9 GENETICS OF FISH POPULATIONS, 200 Robert D. Ward 9.1 Introduction, 200 9.2 Genetic tools, 200 9.3 Statistical tools, 205 9.4 Specimen and species identification, 206 9.5 Fish population genetics, 207 9.6 Genetics of sex determination in fish, 218 9.7 Conclusions, 218 10 BEHAVIOURAL ECOLOGY OF REPRODUCTION IN FISH, 225 Elisabet Forsgren, John D. Reynolds and Anders Berglund 10.1 General introduction, 225 10.2 Introduction to breeding systems, 225 10.3 Parental care, 228 10.4 Sexual selection, 230 10.5 Mating patterns, 236 10.6 Reproductive behaviour and life histories, 238 10.7 Reproductive behaviour and exploitation, 239 10.8 Conclusions, 241 viii Contents Part 3: Fish as Predators and Prey, 249 11 FISH FORAGING AND HABITAT CHOICE: A THEORETICAL PERSPECTIVE, 251 Gary G. Mittelbach 11.1 Introduction, 251 11.2 Foraging behaviour and diet choice, 252 11.3 Foraging models and fish growth, 254 11.4 Feeding rate and group size, 255 11.5 Foraging and habitat selection, 256 11.6 Conclusions, 262 12 FEEDING ECOLOGY OF PISCIVOROUS FISHES, 267 Francis Juanes, Jeffrey A. Buckel and Frederick S. Scharf 12.1 Introduction, 267 12.2 Adaptations for piscivory, 267 12.3 Components of predation, 271 12.4 Prey type and size selectivity, 274 12.5 Predator-size and prey-size relationships, 275 12.6 Population regulation, 277 12.7 Methods of studying predation in the field, 278 12.8 Implications for conservation and management, 279 12.9 Conclusions, 279 13 FISH AS PREY, 284 J. Krause, E.M.A. Hensor and G.D. Ruxton 13.1 Introduction, 284 13.2 Immobility, 285 13.3 Mobility, 287 13.4 Conclusions, 293 Part 4: Fish in Ecosystems, 299 14 TROPHIC ECOLOGY AND THE STRUCTURE OF MARINE FOOD WEBS, 301 Nicholas V.C. Polunin and J.K. Pinnegar 14.1 Introduction, 301 14.2 Food chains and food webs, 302 14.3 Interaction strength in food webs, 312 14.4 Implications of food webs and trophodynamics for fish and fisheries science, 314 14.5 Conclusions, 316 Contents ix 15 COMMUNITY ECOLOGY OF FRESHWATER FISHES, 321 Lennart Persson 15.1 Introduction, 321 15.2 Community patterns and basic ecological performance, 322 15.3 Competition and predation as structuring forces, 325 15.4 Fish community structure, productivity and habitat structure, 328 15.5 Effects of fish on lower trophic components, 331 15.6 From individual-level processes to population dynamics, 334 15.7 Conclusions, 337 16 COMPARATIVE ECOLOGY OF MARINE FISH COMMUNITIES, 341 K. Martha M. Jones, Dean G. Fitzgerald and Peter F. Sale 16.1 Introduction, 341 16.2 Biodiversity, 342 16.3 Habitat associations, 346 16.4 Differences in tropical and temperate production cycles, 350 16.5 Variation in recruitment dynamics, 351 16.6 Conclusions, 352 17 INTERACTIONS BETWEEN FISH, PARASITES AND DISEASE, 359 I. Barber and R. Poulin 17.1 Introduction, 359 17.2 Fish parasite diversity, 360 17.3 Evolution of host–parasite relationships, 365 17.4 Effects of parasites on fish population ecology, 374 17.5 Socioeconomic and human health implications of fish parasites, 376 17.6 Controlling parasite infections, 382 17.7 Recent applications of fish parasitology, 383 17.8 Conclusions, 384 Index, 390 Contributors Geoff Arnold The Centre for Environment, Elisabet Forsgren Department of Marine Fisheries and Aquaculture Science, Lowestoft Ecology, Göteborg University, Kristineberg Ma- Laboratory, Pakefield Road, Lowestoft, Suffolk rine Research Station, SE-450 34 Fiskebäckskil, NR33 0HT, UK. Sweden [email protected] [email protected] The Natural History Museum, I. Barber Edward Llwyd Building, Institute of A.C. Gill Cromwell Road, London SW7 5BD, UK Biological Sciences, The University of Wales [email protected] Aberystwyth, Aberystwyth, SY23 3DA, UK. [email protected] Paul J.B. Hart Department of Biology, University of Leicester, Leicester LE1 7RH, UK Anders Berglund Animal Ecology, Evolution- [email protected] ary Biology Centre, Uppsala University, Norbyvä- gen 18 D, SE-752 36 Uppsala, Sweden E.M.A. Hensor School of Biology, University [email protected] of Leeds, Leeds LS2 9JT, UK [email protected] Ole Brix Department of Zoology, University of Bergen, Allégaten 41, N-5007 Bergen, Norway J.A. Hutchings Department of Biology, [email protected] Dalhousie University, Halifax, NS B3H 4J1, Canada [email protected] Jeffrey A. Buckel Department of Zoology, Center for Marine Sciences and Technology, North Malcolm Jobling The Norwegian College of Carolina State University, 303 College Circle, Fishery Science, University of Tromsø, N-9037 Morehead City, NC 28557, USA Tromsø, Norway [email protected] [email protected] Dean G.